Resistance to herbivory of two populations of <Emphasis Type="Italic">Elodea</Emphasis><Emphasis Type="Italic">canadensis</Emphasis> Michaux and <Emphasis Type="Italic">Elodea nuttallii</Emphasis> Planchon St. John

In this article, we compared the resistance of two introduced populations of Elodea nuttallii and Elodea canadensis to two different herbivores. Samples were collected from the River Rhine and River Rhône in eastern France. The two populations of E. nuttallii differed in their introduction history, whereas E. canadensis was introduced at the same time in the two sites. The Daily Food Consumption (DFC) rates of the two macrophyte populations were evaluated in no-choice experiments using the scraper Lymnaea stagnalis and the shredder Gammarus roeseli. At the same time, we assessed four plant traits: dry matter content (DMC), total nitrogen content, carbon/nitrogen ratio and total phenolic content. The two populations of E. canadensis were consumed at low levels by both the herbivores. L. stagnalis showed a higher DFC on the Rhône population of E. nuttallii than on the Rhine population. No significant difference between the two populations was established with G. roeseli, but the level of DFC was high. This result demonstrates that the assessment of plant palatability should be carried out with several generalist herbivores belonging to various feeding groups (e.g. scrapers or shredders). Although the Rhône population of E. nuttallii had higher levels of phenols than the other populations, it was consistently consumed in greater quantities than E. canadensis. Neither the phenolic contents were not effective against these herbivores, nor the levels of phenolics too low to induce an efficient resistance. The higher DMC and the lower DFC of the two populations of E. canadensis suggest that this introduced plant has co-evolved with indigenous enemies in the introduced range.     

High phenolic content fails to deter mesograzer consumption of <Emphasis Type="Italic">Myriophyllum spicatum</Emphasis> (Eurasian watermilfoil) in New England

Eurasian watermilfoil (Myriophyllum spicatum) is often considered one of the most aggressive macrophyte invaders in freshwater habitats throughout the USA. However, conditions leading to successful milfoil invasions are not well understood. This study sought to illuminate the role of herbivores in determining milfoil invasion success via the potential mechanisms of enemy release and biotic resistance. We determined feeding preferences of three herbivores native to the northeastern United States and measured macrophyte phenolic content, which may act as an herbivore feeding deterrent. We found that phenolic content in milfoil was more than two times higher than in the most abundant native macrophytes at our study sites, consistent with enemy release. However, laboratory feeding experiments demonstrated that milfoil phenolics did not deter amphipod (Hyalella azteca), snail (Physella sp.), or weevil (Euhrychiopsis lecontei) herbivory. Furthermore, amphipod consumption rates in our study were an order of magnitude higher than amphipod consumption rates reported in milfoil’s native range, contrary to the predictions of enemy release. Amphipods and snails from habitats invaded by milfoil consumed similar quantities of both milfoil and the low-phenolic native plant Elodea canadensis. In contrast, weevils consumed milfoil but not E. canadensis in choice experiments. Amphipods collected from milfoil-free habitats also readily consumed milfoil, and they consumed 2.5 times more milfoil than E. canadensis in a choice feeding trial. These results suggest that high phenolic levels do not prevent native herbivores from consuming invasive milfoil. Instead, native generalist grazers like amphipods and snails may limit milfoil proliferation and provide a measure of biotic resistance.     

Consortium of Higher Aquatic Plants and Microalgae Designed to Purify Sewage of Heavy Metal Ions

We selected higher aquatic plants (HAP) and microalgae possessing a high sorption capacity in respect to heavy metals to form a consortium designed to purify contaminated aquatic ecosystems. Accumulation of heavy metals Cd²⁺, Cu²⁺, Pb²⁺, and Zn²⁺ was investigated in plants Pistia stratiotes, Elodea canadensis, and Lemna minor and green microalgae Chlorella vulgaris ВВ-2, Ankistrodesmus sp. ВI-1, Chlamydomonas reinhardtii В-4, and Scеnеdеsmus quadricauda В-1. It was found that intense accumulation of metals occurs in cultures of HAP Pistia stratiotes and Elodea canadensis. These plants are macroconcentrators of zinc, lead, and copper and microconcentrators of cadmium. Out of the examined cultures of microalgae, effective bioaccumulators of heavy metals were C. vulgaris ВВ-2 and Ankistrodesmus sp. ВI-1. It was shown that heavy metals are selectively taken up from the medium in the series Zn²⁺ > Cu²⁺ > Cd²⁺ > Pb²⁺. In order to produce a consortium of higher aquatic plants and microalgae for purification of polluted aquatic ecosystems, we investigated interaction of HAP P. stratiotes and E. canadensis with microalgae C. vulgaris ВВ-2 and Ankistrodesmus sp. ВI-1 in the course of their cocultivation. Neutral relations were detected between the cells of microalgae C. vulgaris ВВ-2 and Ankistrodesmus sp. ВI-1 and HAP E. canadensis. At the same time, the cells of Ankistrodesmus sp. ВI-1 and HAP P. stratiotes formed a symbiosis. Microscopic examination showed numerous points where the cells of microalgae Ankistrodesmus sp. ВI-1 were attached to the roots of P. stratiotes plants. We tested an opportunity to employ the association between P. stratiotes and Ankistrodesmus sp. ВI-1 for purification of simulated wastewater polluted with heavy metal ions. This consortium proved to be capable of eliminating contaminants from the sewage, reducing their level in the sewage to standard values, and active accumulation of heavy metal ions.     

Diversity and composition of bacterial community in the rhizosphere sediments of submerged macrophytes revealed by 454 pyrosequencing

Freshwater lake sediments support a variety of submerged macrophytes that may host groups of bacteria exerting important ecological functions. We collected three kinds of commonly found submerged macrophyte species (Ceratophyllum demersum, Vallisneria spiralis and Elodea nuttallii) to investigate the bacterial community associated with their rhizosphere sediments. High-throughput 454 pyrosequencing and bioinformatics analyses were performed to examine the diversity and composition of the bacterial community. The results obtained indicated that the diversity of the bacterial community associated with the rhizosphere sediments of submerged macrophytes was significantly lower than that of the bulk sediment. Remarkable differences in the bacterial community composition between the rhizosphere and bulk sediments were also observed.     

Inter-species variation in unpalatability does not explain polymorphism in a mimetic species

Conspicuous colouration in unpalatable organisms acts as a warning signal of their unprofitability, a phenomenon known as aposematism. The protection conferred by such colouration can lead to evolutionary convergence in warning signals between aposematic species, because sharing warning signals reduces the per capita cost of predator learning. Consequently, most aposematic species display a single colour pattern and participate in a single mimetic community (i.e. mimicry ring) at any given locality. However, some, like the Amazonian butterfly Heliconius numata, are polymorphic and participate in several mimicry rings within the same locality. We tested whether the unexpected polymorphism of H. numata could be due to a weak defence against predators. Poorly defended species participating in a mimicry ring are subject to negative frequency dependent selection, because their presence weakens the protection provided by the shared signal. This could promote polymorphism and participation in multiple mimicry rings. Using wild caught great tits (Parus major), we compared the palatability of H. numata to one of its locally monomorphic co-mimics (Mechanitis polymnia) and to two other locally monomorphic Heliconius species (H. melpomene and H. erato). The tested birds strongly rejected the polymorphic species H. numata, as well as the two other Heliconius species. Unexpectedly, a significantly weaker rejection was found towards M. polymnia, which relies on different toxic compounds to Heliconius. Our study demonstrates that the origin of polymorphic mimicry in H. numata is unlikely to stem from low unpalatability and raises new questions on defence variation within mimetic communities.     

Patterns of microbial food webs in Mediterranean shallow lakes with contrasting nutrient levels and predation pressures

Hygraula nitens is a New Zealand native moth with aquatic larvae that feed on submerged aquatic plants. The larvae have been mainly observed using native Potamogeton and Myriophyllum species as a food source, although some studies reported larvae feeding on the alien macrophytes Hydrilla verticillata, Lagarosiphon major and Ceratophyllum demersum. Experimental mesocosm studies showed larvae had a major effect on H. verticillata, C. demersum, L. major, Elodea canadensis and Egeria densa. In both no choice and choice experiments H. nitens larvae showed a clear preference for and the highest consumption of C. demersum, while the native macrophyte Myriophyllum triphyllum ranked fourth out of five alien and two native plant species, indicating a preference of the larvae for alien macrophytes. Additional choice experiments using C. demersum, sampled from different waters in NZ, illustrated that there was a clear difference in H. nitens preference for plants based on their source. However although C. demersum had the lowest leaf dry matter content (LDMC) compared with the other macrophytes, neither the LDMC nor leaf carbon, nitrogen, phosphorus or total phenolic contents alone could explain the preferences of H. nitens, and we conclude that food choice is based on a combination of these and/or additional factors.     

Differences in herbivore damage and performance among <Emphasis Type="Italic">Arctium minus</Emphasis> (burdock) genotypes sampled from a geographic gradient: a common garden experiment

Performance of plant species does not necessarily decline as they approach their geographic range limits. One reason for this may be a loss of natural enemies in marginal populations. Such patterns have been found in native species, but also may occur for exotics if they have not already escaped their herbivores in invaded regions. For instance, the Eurasian biennial Arctium minus (common burdock) is attacked by a variety of native and introduced insects in its new North American range. Previously, research has shown that damage by these herbivores strongly decreases towards the northern range limit of this species. This gradient might reflect a genetic cline in resistance to herbivores, or geographic variation in herbivore abundance. To distinguish between these possibilities, herbivore damage to leaves and seeds of A. minus was measured in a common garden experiment with genotypes sampled from 11 populations along a 550 km transect extending from southern Ontario towards burdock’s northern range limit. As well, a freezing tolerance experiment was performed with the important lepidopteran seed predator Metzneria lappella, and palatability experiments were performed with two generalists, the snail Cepaea nemoralis and the moth Trichoplusia ni. Although there were some differences in damage among populations, results indicated that latitudinal differences in herbivore damage are not explained by genotypic differences among populations, but instead are likely to result from the absence of herbivores from colder sites. Escape of A. minus from its usual herbivores may increase performance of marginal populations, and contribute to future spread.     

Molecular evidence for hybridization between invasive <Emphasis Type="Italic">Solidago canadensis</Emphasis> and native <Emphasis Type="Italic">S</Emphasis>. <Emphasis Type="Italic">virgaurea</Emphasis>

Hybridization between alien and native species is biologically very important and could lead to genetic erosion of native taxa. Solidago × niederederi was discovered over a century ago in Austria and described by Khek as a natural hybrid between the alien (nowadays regarded also as invasive) S. canadensis and native S. virgaurea. Although interspecific hybridization in the genus Solidago is considered to be relatively common, hybrid nature of S. × niederederi has not been independently proven using molecular tools, to date. Because proper identification of the parentage for the hybrid Solidago individuals solely based on morphological features can be misleading, in this paper we report an additive polymorphism pattern expressed in the ITS sequences obtained from individuals representing S. × niederederi, and confirm the previous hypothesis that the parental species of this hybrid are S. canadensis and S. virgaurea. Additionally, based on variability at the cpDNA rpl32-trnL locus, we showed that in natural populations hybridization occurs in both directions.     

Can snail herbivory influence the outcome of competition between Elodea species?

Elodea nuttallii is more competitive than Elodea canadensis but is also more palatable to herbivores. We tested if grazing by generalist invertebrate herbivores could modify the competitive abilities of E. nuttallii, and thereby influence the outcome of the competition between the two Elodea species. The influence of snail herbivory on the competitive interactions between the two Elodea species cultivated together in indoor tanks was tested for 2 months. The presence of Lymnaea stagnalis in the tanks reduced significantly the final number of rootings, as well as the final biomass of E. nuttallii, but not the surface colonised by this species, whereas only the final number of rootings of E. canadensis was reduced. However, E. nuttallii remained the dominant species. We conclude that snail herbivory can influence the outcome of competition between Elodea species. However, snails are probably not an effective biological control agent of E. nuttallii.     

Deer feeding selectivity for invasive plants

Native generalist herbivores might limit plant invasion by consuming invading plants or enhance plant invasion by selectively avoiding them. The role of herbivores in plant invasion has been investigated in relation to plant native/introduced status, however, a knowledge gap exists about whether food selection occurs according to native/introduced status or to species. We tested preference of the native herbivore white-tailed deer (Odocoileus virginianus) for widespread and frequently occurring invasive introduced and native plants in the northeastern United States. Multiple-choice deer preference trials were conducted for the species and relative preference was determined using biomass consumption and feeding behavior. While more native than introduced plant biomass was consumed overall, deer food selection varied strongly by plant species. Results show consistent deer avoidance of several invasive introduced plants (Alliaria petiolata, Berberis thunbergii, and Microstegium vimineum) and a native plant (Dennstaedtia punctilobula). Other invasive introduced plants (Celastrus orbiculatus, Ligustrum vulgare, and Lonicera morrowii) and a native plant (Acer rubrum) were highly preferred. These results provide evidence that herbivore impacts on plant invaders depend on plant species palatability. Consequently, herbivore selectivity likely plays an important role in the invasion process. To the extent that herbivory impacts population demographics, these results suggest that native generalist herbivores promote enemy release of some plant invaders by avoiding them and contribute to biotic resistance of others by consuming them.     

Response of invasive macrophyte species to drawdown: The case of Elodea sp.

The response to drawdown of vegetative fragments (whole plants, shoot fragments and turions) of two invasive macrophyte species, Elodea canadensis and Elodea nuttallii, was studied through laboratory experiments. In addition, field observations were made on the colonisation of a wetland by E. nuttallii before and after a natural drawdown. The survival and the growth of vegetative fragments of E. nuttallii were higher than those of E. canadensis after an artificial drawdown of several days. In the field the recolonisation by E. nuttallii of a wetland that was drained for 10 weeks during a summer drawdown was very rapid, the abundance of this macrophyte species being not affected by the drawdown event. We conclude that E. nuttallii possesses a high resilience to desiccation and that a summer drawdown would not be efficient in the control of this invasive species.     

Long-term adaptation of <Emphasis Type="Italic">Escherichia coli</Emphasis> to methanogenic co-culture enhanced succinate production from crude glycerol

Escherichia coli can hardly grow anaerobically on glycerol without exogenous electron acceptor. The formate-consuming methanogen Methanobacterium formicicum plays a role as a living electron acceptor in glycerol fermentation of E. coli. Wild-type and mutant E. coli strains were screened for succinate production using glycerol in a co-culture with M. formicicum. Subsequently, E. coli was adapted to glycerol fermentation over 39 rounds (273 days) by successive co-culture with M. formicicum. The adapted E. coli (19.9 mM) produced twice as much succinate as non-adapted E. coli (9.7 mM) and 62% more methane. This study demonstrated improved succinate production from waste glycerol using an adapted wild-type strain of E. coli with wild-type M. formicicum, which is more useful than genetically modified strains. Crude glycerol, an economical feedstock, was used for the cultivation. Furthermore, the increase in methane production by M. formicicum during co-culture with adapted E. coli illustrated the possibility of energy-saving effects for the fermentation process.     

Response of invasive macrophyte species to drawdown: The case of Elodea sp.

The response to drawdown of vegetative fragments (whole plants, shoot fragments and turions) of two invasive macrophyte species, Elodea canadensis and Elodea nuttallii, was studied through laboratory experiments. In addition, field observations were made on the colonisation of a wetland by E. nuttallii before and after a natural drawdown. The survival and the growth of vegetative fragments of E. nuttallii were higher than those of E. canadensis after an artificial drawdown of several days. In the field the recolonisation by E. nuttallii of a wetland that was drained for 10 weeks during a summer drawdown was very rapid, the abundance of this macrophyte species being not affected by the drawdown event. We conclude that E. nuttallii possesses a high resilience to desiccation and that a summer drawdown would not be efficient in the control of this invasive species.     

Influence of Aquatic Plants of Different Ecological Groups on Zooplankton Distribution and Abundance

The distribution of zooplankton has been studied in a waterbody whose entire area is occupied by phytocenoses with the dominance of plants of different ecological groups. It is shown that, in the phytocoenosis of the submerged hydrophyte Elodea canadensis, zooplankton is involved in the classical grazing trophic web. The zooplankton community in this phytocoenosis is mainly made up by algaephagous species. In the habitats occupied by emergent plant Carex riparia, zooplankton is more closely associated with the components of the “microbial loop,” where the most abundant groups are predators and bacterio-detritophagous species.     

Seasonal variations in moisture use in a piñon–juniper woodland

In water-limited environments of the intermountain region of North America, summer precipitation may play a role in the structure and function of aridland communities and ecosystems. This study examined the potential reliance on summer precipitation of two widespread, coexisting woody species in the southwestern United States, Pinus edulis Englmn. (Colorado piñon) and Juniperus osteosperma (Torr) Little (Utah juniper). The current distributions of P. edulis and J. osteosperma are highly suggestive of different dependencies on summer rainfall. We hypothesized that P. edulis was dependent on summer precipitation, utilizing summer precipitation even during extremely dry summers, whereas J. osteosperma was not dependent, using summer precipitation only when amounts were above some minimum threshold. Using sap flux and stable isotopic methods to assess seasonal water sources and water use efficiency, we examined the response of these two species to seasonal variations in moisture at a site located near the northern limits of the North American monsoon. Both sap flux and isotopic results indicated that P. edulis was responsive to summer rain, while J. osteosperma was not. Following summer rain events, sap flux density increased in P. edulis for several days, but not in J. osteosperma. Isotopic evidence indicated that P. edulis took up summer-derived moisture to a greater extent than J. osteosperma. Values of the natural abundance stable isotope ratio of carbon of leaf soluble carbohydrates increased over the summer for P. edulis, indicative of assimilation at higher water use efficiency, but were invariant for J. osteosperma. Our results supported the hypothesis that P. edulis and J. osteosperma are differentially sensitive to summer precipitation and are discussed in the light of potential changes in the seasonality of precipitation associated with climate change.     

Spatio-temporal variation of strawberry aphid populations and their parasitoids

Aphids are common herbivores in the strawberry crop that can reduce plant vigor and fruit quality and also transmit viruses. Aphid species prefer diverse plant organs, which represent particular habitats of different quality for aphids and for the development of natural enemies’ populations. Different habitat units (young leaves, mature leaves, buds, flowers) of strawberry were sampled fortnightly during all seasons. We identified seven aphid species, namely Chaetosiphon fragaefolii, Aphis gossypii, and Macrosiphum euphorbiae, the most abundant. During the autumn, C. fragaefolli and M. euphorbiae were scarce and A. gossypii was denser on mature leaves, while during summer M. euphorbiae was absent. During the winter, C. fragaefolii predominated on buds and young leaves, A. gossypii on flowers, and both species on mature leaves. During the spring, C. fragaefolii was even more abundant on buds, A. gossypii predominated on mature leaves, and the three species were equally abundant on flowers and young leaves. Parasitoids emerged from A. gossypii, M. euphorbiae and Myzus persicae, but not from C. fragaefolii. Three Aphidius and two Aphelinus species were recovered. All primary parasitoid species emerged from A. gossypii, and secondary parasitoids emerged only from this aphid. Aphis gossypii parasitism on mature leaves was markedly higher in winter and summer than in autumn and spring. Parasitism of A. gossypii was independent of its density, and the number of parasitized aphids was never higher than six. Our results contribute to define the most appropriate sample unit to estimate aphid density of different species and provide information about seasonal natural parasitism.     

Invasive alien plants of Russia: insights from regional inventories

Recent research on plant invasions indicates that some parts of the world are understudied with temperate Asia among them. To contribute towards closing this gap, we provide a standardized list of invasive alien plant species with their distributions in 45 Russian regions, and relate the variation in their richness to climate, socioeconomic parameters and human influence. In total, we report 354 invasive alien species. There are, on average, 27 ± 17 (mean ± SD) invasive plants per region, and the invasive species richness varies from zero in Karelia to 71 in Kaluga. In the European part of Russia, there are 277 invasive species in total, in Siberia 70, and in the Far East 79. The most widespread invaders are, in terms of the number of regions from which they are reported, Acer negundo, Echinocystis lobata (recorded in 34 regions), Erigeron canadensis and Elodea canadensis (recorded in 30 regions). Most invasive species in Russia originate from other parts of temperate Asia and Europe. There were significant differences in the representation of life forms between the European, Siberian and Far East biogeographical regions, with perennials being over-represented in the Far East, and shrubs in the European part of Russia. The richness of invasive species can be explained by climatic factors, human population density and the percentage of urban population in a region. This publication and the associated dataset is the first comprehensive treatment of the invasive flora of Russia using standardized criteria and covering 83% of the territory of this country.     

Seasonal variability in the palatability of freshwater macrophytes: a case study

The pond snail Lymnaea stagnalis (L.) was used for a laboratory assessment of seasonal variation in palatability of three freshwater macrophytes: Potamogeton lucens, Elodea canadensis and E. nuttallii. For each species, 2–5 populations were investigated in spring and in summer. Preliminary results showed that the feeding rate of similarly-aged snails bred under standard conditions was stable over time. In contrast, snail feeding rate on the three macrophyte species decreased from spring to summer, which was therefore interpreted as a decrease in plant palatability. This decrease was probably due to tissue maturation, as suggested by the concomitant increase in the dry matter content of leaves of the three species. The high palatability of the species studied during the spring may prove detrimental in cases of strong herbivore pressure, and could have consequences for macrophyte distribution among aquatic habitats.     

An invasive plant provides refuge to native plant species in an intensely grazed ecosystem

Invasion by exotic plant species and herbivory can individually alter native plant species diversity, but their interactive effects in structuring native plant communities remain little studied. Many exotic plant species escape from their co-evolved specialized herbivores in their native range (in accordance with the enemy release hypothesis). When these invasive plants are relatively unpalatable, they may act as nurse plants by reducing herbivore damage on co-occurring native plants, thereby structuring native plant communities. However, the potential for unpalatable invasive plants to structure native plant communities has been little investigated. Here, we tested whether presence of an unpalatable exotic invader Opuntia ficus-indica was associated with the structure of native plant communities in an ecosystem with a long history of grazing by ungulate herbivores. Along 17 transects (each 1000 m long), we conducted a native vegetation survey in paired invaded and uninvaded plots. Plots that harboured O. ficus-indica had higher native plant species richness and Shannon–Wiener diversity H′ than uninvaded plots. However, mean species evenness J was similar between invaded and uninvaded plots. There was no significant correlation between native plant diversity and percentage plot cover by O. ficus-indica. Presence of O. ficus-indica was associated with a compositional change in native community assemblages between paired invaded and uninvaded plots. Although these results are only correlative, they suggest that unpalatable exotic plants may play an important ecological role as refugia for maintenance of native plant diversity in intensely grazed ecosystems.